Lawrence Alexander Sidney (Lawrie) Johnson was a taxonomic botanist notable for the outstanding breadth of his interests and expertise, the rigour of his scientific approach, and the intensity with which he defended scientific conclusions and opinions. His major contributions came through broad synthesis so that systematic studies were integrated with evolutionary and ecological considerations. In a field often characterized by solitary workers, his investigations mostly involved colleagues, and he encouraged them to share his wide-ranging outlook while he also relied on their efforts. With colleagues he tackled many large plant groups, often delimiting taxa and resolving questions of relationships far beyond the extent that could be published in his lifetime: jointly authored publications continued to appear for some years after his death, since Ken Hill, Karen Wilson and I acknowledged that he had a substantial role in work still being finalized by each of us.
Johnson's achievements were recognized by many awards, to an extent remarkable for a scientist in the field of systematic botany and attached to one institution, the Royal Botanic Gardens Sydney, for the whole of his professional career, as Botanist (1948-72), Director (1972-85) and Honorary Research Associate (1986-97).
Since Johnson was well known for his strongly held views, passionately expressed, it was in character that, when he knew that he was terminally ill, he prepared the following statement to be read at his funeral. It sums up much of his life, with characteristic messages for those who heard it:
I have never been inclined to follow convention, and therefore will not be silent even at my own funeral. The first thing I have to say is in all sincerity to express my thanks for many opportunities to lead a satisfying life which, though not as long as I would have hoped, has been long enough for me to feel very fairly treated. These opportunities began with my deeply respected and loved parents, Sid and Emily Johnson, and my two sisters Valerie and Nancy. Teachers, especially my science teachers at school, Mr Clark and Mr Roberts, enabled me to perceive clearly in adolescence the absurdity of all forms of superstition and fanciful belief. They were followed at the University of Sydney by some outstanding teachers at that level, especially Eric Ashby (later Lord Ashby) and Sir Rutherford Robertson. These men showed me that botany was indeed a science, and I abandoned my first love, chemistry, to follow botany, although in a field very different from those of Ashby and Rutherford [Robertson].
In my first year botany lab I had the great good fortune to meet Merle, who has been the epitome of the word steadfastness throughout our subsequent lives. Our five children, Chris, Sylvia, Nicholas, Quentin and Sandy [Alexander], have borne with us, and we with them for up to 45 years, and have been a great support during my final illness. I love them all, though they may not always believe it.
Our personal friends have helped us in our lives for many years, and are immensely valued by us. On the professional side, I pay tribute to all my predecessors as Directors of the Royal Botanic Gardens, with a special tribute to Robert Henry Anderson who encouraged me to join the Sydney Botanic Gardens, and who was, as a canny Scot, a friend as well as a mentor at all times. Among many others at the RBG, my very special mention [and] appreciation must go to Dr Barbara Briggs who has been my close research and administrative colleague, and the very best personal friend since 1960. You will note that I do not say 'managerial colleague' – I have found that administrators, though they can be beastly, are not as manifestly so [as] those who call themselves managers.
Good friends and stimulating colleagues have been many throughout Australia and the world, and I am indeed sorry that I cannot continue to join in research with them. I conclude with two more saddening observations. First, the RBG has been abominably treated by politicians and botanically totally ignorant managers over the past year. This was not formerly the case and, in the days of such enlightened politicians as Neville Wran, the right sort of people were allowed to pursue the proper aims and objects of a botanic gardens. Second, a larger manifestation of these tendencies is a worldwide concentration on managerialism, a superficial approach and indeed a cruel perversion of sensible and decent management.
I have not seen an effective countering of this approach, but I can safely say that I have never met a decent managerialist yet. I hope, indeed, that some of you will live to see the change that must come.
I hope that I have not offended too many people in saying that since my eyes were opened in adolescence I have seen no reason to need religious belief, and particularly the notion of continuation of 'life' after death. I have been happy to accept death as the natural end of life, which treats some of us well, and some of us ill. It has treated me well.
I thank you all for coming and wish you satisfaction in your lives.(1)
Born in the Sydney suburb of Cheltenham on 26 June 1925, Lawrie was the third child of Algernon Sidney (Sid) Johnson, an accountant in the hospitals section of the New South Wales Public Service and Emily Margaret Johnson (née Manson), with older sisters Valerie and Nancy. He was early recognised as a bright child, attending Beecroft Public School and Parramatta High School. His father died when he was nine years old but his mother encouraged his academic abilities and, since early teenage years, he planned to study science, keen to understand and classify features of the world around him. Throughout his life he expressed gratitude to teachers whom he respected and who fostered his academic interests. He enjoyed the bushland then persisting around Cheltenham and Devlins Creek, but his early interest focused on chemistry and he was equal first in the State in that subject at the Leaving Certificate examination in 1941.
Enrolling at the University of Sydney, having been awarded a scholarship, he took Botany with no intention of continuing in that discipline, but the lively minds in the Botany Department at that time attracted him to the subject. He was fortunate there to meet Professor Eric Ashby (later Lord Ashby), plant physiologist Bob (later Sir Rutherford) Robertson, ecologist Noel Beadle and geneticist Newton Barber. This was a time of wide-ranging, indeed re-awakening, interest in Australia's flora and vegetation, and botanists were searching for understanding of processes of evolution and variation, seen in terms of ecology and physiology. In his final undergraduate research year he began a taxonomic revision of Casuarina (Casuarinaceae) under the supervision of Patrick Brough, but with advice and guidance from R.H. (Bob) Anderson and Joyce Vickery at the National Herbarium of New South Wales at Sydney's Royal Botanic Gardens. The project proved a far larger undertaking than his supervisor ever imagined and studies in Casuarinaceae continued among his interests throughout his career. In 1971 he was awarded a DSc by the University of Sydney.
Graduating with First Class Honours in 1948, he joined the staff of the National Herbarium in Sydney. The Herbarium was the scientific arm of the Royal Botanic Gardens, administered as part of the New South Wales Department of Agriculture. During the directorship of Joseph Henry Maiden (1896-1924) the Herbarium had been founded, developed and linked to other organizations world-wide through Maiden's extensive correspondence and one visit to Britain. Maiden had been extraordinarily energetic with programmes of community lectures and taking a leading part in Sydney's emerging scientific associations. As a result the Gardens and its science had held high profile and esteem.(2) This had sadly deteriorated over subsequent decades so that Johnson joined a small group of botanists there at a time when taxonomy was unfashionable and Sydney's herbarium seemed something of a scientific backwater. An injection of young staff was, however, already starting a renewal after a less productive period.
Johnson realized the importance of collaboration with scientists elsewhere and soon developed close links with botanist colleagues Spencer Smith-White (later professor of genetics at the University of Sydney) and J.L. (Jack) Willis at the Museum of Applied Arts and Sciences (forerunner of the Powerhouse Museum) in Sydney, as well as phytochemist Howard McKern. Soon an even more productive collaboration was established with Lindsay Pryor, then Director of Parks and Gardens in the Australian Capital Territory, later foundation professor of botany at the Australian National University.
At the Herbarium he took over curatorial management of a wide range of plant groups. He and his colleagues set about improving the botanical order and information content of the collection, critically assessing the classification of many plant groups and the identification of the specimens. Through such work with the specimen collections, field studies and providing identifications for enquirers, Johnson acquired a remarkably wide knowledge of vascular plants in Australia and the rest of the world, knowledge he generously shared with others. Always he was intensely aware that the specimen collections were exemplars of the populations in nature; for this reason he sorted the collections geographically in detail, so that they would reveal trends and discontinuities in the species' distribution.
Since a responsibility of the Herbarium was to provide sources of information on the plants of New South Wales, a Flora of the State was begun, issued in segments from 1961 to 1984 (Contributions from the National Herbarium of N.S.W.-Flora Series). With hindsight this was a misguided project, scholarly and authoritative but so detailed that progress was painfully slow; too technical for non-specialist users but lacking some features of full taxonomic revisions. Good work was done for the project but progress was almost in spite of the concept of the series. Johnson's treatment of the cycads [13] was one of the first and most thorough of the series but, characteristically, he accompanied it by a more notable work on cycads world-wide [12].
Johnson's publication list and reputation among systematic botanists grew and he was in 1962 appointed as Australian Botanical Liaison Officer, a twelve-month posting to the Royal Botanic Gardens, Kew. He was a member of the Flora of Australia Committee of ANZAAS and had been part of a delegation in 1960 to the Prime Minister's Department seeking support for a national Flora project. He was later appointed to the committee that eventually led to success in that aim, the Interim Council of the Australian Biological Resources Study. During 1973-5 that Council set guidelines, helped to make clear to the Australian Government the great need for the Flora of Australia, and visited institutions in major centres to discuss biologists' needs and to promote the new projects envisaged. The Flora programme began to receive Australian Government funding in 1979 and has been acknowledged internationally as highly effective, its success partly based on the associated grants programme and the establishment of a core of editorial staff overseeing the work.
In 1968 Johnson had been appointed as Deputy Chief Botanist, the deputy to the Botanic Gardens Director, but when Knowles Mair retired in 1970, Johnson did not seek the senior position. He had seen directors of the Gardens submerged in administrative and horticultural detail and it was only later, when he acted briefly in the position, that he realized how the directorship could be handled differently. He saw that change was possible with a clear concept of the role of the organization and a vision for its future [82]. When the position again became vacant in 1972 he was appointed to it.
Johnson was fortunate that, during much of his thirteen years as Director, Neville Wran was Premier of New South Wales. Johnson sought any opportunities to interest Wran in the Gardens. For his part, Wran still remembered his own role as counsel assisting Lionel Murphy, barrister for the Public Service Association, in a case brought by that union before the New South Wales Industrial Commission many years earlier. That case reviewed salaries of Scientific Officers in the New South Wales Public Service, and Gardens' botanists had taken a prominent role. In 1980 Wran transferred the Gardens into the Premier's Department administration, to join with museums and other cultural organizations. The Gardens was Australia's oldest continuing scientific institution, predating modern administrative structures but, since the establishment of the New South Wales Department of Agriculture, had been administered as one of its agencies. It had been seen as peripheral to Agriculture's role but was instantly 'at home' among the diverse cultural agencies.
Johnson grasped the new opportunities with energy. The Royal Botanic Gardens and Domain Trust was set up, government funding was provided for a new building to house the herbarium and scientific programmes, freeing much-needed space for education programmes, visitor services and the necessarily increasing administrative staff. Community support was fostered and channelled into an active 'Friends of the Royal Botanic Gardens' organization. An education programme for school students was initiated and, with the start of volunteer programmes, there were guided tours of the Gardens and exhibitions on botanical and biodiversity conservation themes. The first of the Tropical Centre glasshouses (the Pyramid) was completed and a larger glasshouse (the Arc) planned and funded. Preparation for the XIII International Botanical Congress in Sydney in 1981 was another commitment for much of the scientific staff.
Funding was obtained for the development of a satellite garden at Mount Tomah on the escarpment of the Blue Mountains as a State-Commonwealth bilateral programme, associated with Australia's Bicentennial celebrations. Also funded was the development of the Mount Annan Garden in Sydney's south-west, to display and conserve Australian native plant species. The Mount Tomah Garden site had been donated to the State by the Brunet family in the 1960s, to be part of the Gardens' organization, but its development had languished with few resources for development. The Mount Annan site was land previously resumed by the State Government for its scenic and open-space value and selected by the Gardens staff as highly suitable for an Australian plant garden. The documentation and vouchering of plant accessions at Mount Annan set new standards in living collections databases. Both new gardens were opened after Johnson's retirement, as part of the Bicentennial celebrations, Tomah in late 1987 and Annan in 1988; but the main elements of planning, construction and planting had been put in place in his time.
In contrast to this expansion, Johnson encouraged the Government to set up a separate administration for Sydney's large Centennial Park, hitherto part of the Gardens' responsibilities. He considered its largely recreational and open-space functions as very different from the botanical and educational roles of a botanic garden. He did, however, value Sydney's Domain as a vital buffer that surrounded the Gardens on three sides and he argued strongly against any proposed encroachments on it. With the advent of Sydney's Summer Festival, and despite some initial reluctance, he welcomed the increasing use of the Domain for open-air art exhibitions and large events such as opera and symphony concerts.
A new project was initiated, with his enthusiastic encouragement, to produce a Flora of New South Wales, in a far more user-friendly style than the earlier Flora Series. Gwen Harden was appointed as editor and project leader and the Flora was produced in four large volumes in successive years from 1990 to 1993, making information and means of identification available for more than 6,000 species of native and naturalized plants of the State.(3) Johnson was not so positive about the development of computerized databases for the herbarium collection and living plant collections data, but he accepted his staff's advice and these were established. He strongly supported women horticulturalists joining the Gardens' staff, entering an area that – unlike the scientific section – had been an all-male preserve.
Such a rapid pace of development, especially in the latter years of his directorship, transformed the organization, but the resources of funding and especially of staff were stretched thin. As a result, he regretted that there was some reduction in horticultural standards in the Gardens in Sydney, but he hoped that a period of consolidation and remedying of such problems would follow this developmental phase. Always he spoke most enthusiastically about the scientific role of the Royal Botanic Gardens, but much of his legacy there is in its new satellite gardens and community outreach.
During this time he did not abandon research but relied increasingly on co-operation with colleagues. In other cases he encouraged staff botanists to take over work in plant groups with which he had previously been concerned. He continued to be interested in Proteaceae, Myrtaceae, Cyperaceae, cycadophytes and the genus Grevillea, although colleagues Peter Weston, Peter Wilson, Karen Wilson, Ken Hill and Don McGillivray respectively took up research and established their positions as authorities in these groups, building on his earlier work in each case.
On 18 November 1950 Lawrie married Merle Margaret Asta Hodge whom he had met as a first year Botany student, although Merle did not continue with her course. They subsequently lived mostly at Northbridge, a northern Sydney suburb. Merle was an outstanding support in Lawrie's career, typing many of his earlier manuscripts and tolerating his long hours at the office, especially when he maintained his research programmes despite heavy administrative responsibilities. Her support for his career was most dramatically shown when the opportunity came for him in 1962 to take the position of Australian Botanical Liaison Officer at the Royal Botanic Gardens, Kew, London, involving an absence from Sydney of fifteen months. The miserly policy of the New South Wales Government at the time did not provide adequate travelling support for dependants, although this was a man with wife and four, soon to be five, children. Merle steadfastly insisted that he should take the position while she and the children remained in Sydney. At that time they did not know that Merle would later work in an airline administration, earning opportunities for much travel by all the family. It was Merle's outstanding hospitality and concern for people that kept together a wide circle of friends, especially keeping in touch with friends from school and university days.
When Lawrie's long working hours were set aside for time with their children, Christopher, Sylvia, Nicholas, Quentin and Alexander (Sandy), they often jointly constructed magnificent large creations of meccano or set up model trains that ran on lines that looped from room to room in the house for days. At other times all the family, or one of the children with their father, went travelling back-roads and camping in the bush, rarely turning for home until dusk on the last day. There were also family summer holidays near the beach at Hat Head or South West Rocks on the north coast of New South Wales and Saturday mornings watching the sons playing soccer. Chris, Nicholas and Sandy later worked in various aspects of information technology, Chris at the University of Technology, Sydney. Quentin became a fire fighter in the New South Wales fire brigades. Sylvia's career in materials science, with a focus on ceramics, took her to California and distinguished roles at the Stanford Research Institute and NASA.
Johnson's energy and achievements did not come from an especially robust constitution. Through much of his career, Johnson suffered from an anxiety condition and his ability to sustain such a level of activity was partly a credit to modern pharmaceuticals. A whiplash neck injury resulted from a car accident while returning from a scientific conference; this frequently gave trouble but did not deter him from active fieldwork. For many years he also suffered from a persistent stomach ulcer. Earlier, during his first year as an undergraduate, a diagnosis of tuberculosis, which more severely affected his sister Nancy, had interrupted his studies. That enforced break, until he resumed his course a year later, was used to read a wide range of literature. Years later he observed that this interruption had been beneficial in the long term, broadening his knowledge in a way that would have been unlikely in the midst of studies or the demands of a career. Aided by secretaries and colleagues, through much of his career, including his busy years as Director, he managed to 'protect' time for a brief siesta after lunch on most days. Then he would customarily work far into the evening.
Apart from botany, one of Johnson's long-standing interests was in comparative languages. The processes of development, modification and migration of languages revealed many fascinating similarities to processes of biological evolution. Respect for other languages, for him, included correct pronunciation and he persevered in trying to correct the pronunciation of foreign words and botanical names by colleagues. A guide to the pronunciation of botanical names was one outcome [110]. He studied Russian at evening courses and took great interest in languages wherever he travelled. He frequently kept his colleagues amused with limericks and other verses.
Another interest fostered by evening courses was mathematics, at the time when newer mathematical concepts were beginning to be taught in schools. For some years in mid-career he took a serious interest in number theory and topology, finding that they involved satisfyingly elegant concepts. These studies stimulated him to analyse the theoretical basis of the numerical pheneticists in the 1960s and to publish his comprehensive rejection of that approach [32].
Other lifelong interests – all actively shared with Merle – were in classical music, railways (an interest dating from school days) especially historic trains, tennis with his family and younger friends, travel, and good food and wine. Visits or excursions with his grandchildren, six in the Sydney region and two in California, were also much enjoyed in later years.
Johnson did not intend to retire in 1985 but Government policies of that time strongly favoured youth employment and he found that retirement at age 60 years (rather than the expected 65) was demanded. This was in sharp contrast to policies opposed to age-discrimination in employment implemented only a few years later. Retiring when many developments that he had started were yet to come to fruition was a bitter matter, and this coloured his view for some years.
On retirement he was appointed by the Royal Botanic Gardens Trust as Director Emeritus and as an Honorary Research Associate; facilities for research were provided. He continued to keep better informed on current scientific literature than many much younger botanists and was a source of vast knowledge and help to young staff members at the Herbarium. He greatly welcomed the development of a DNA laboratory for molecular plant systematics at the Royal Botanic Gardens and actively followed literature in this important field. He recognized that macromolecular systematics was now the most important source of new phylogenetic understanding. In conjunction with critically assessed morphology and phylogenetic reasoning, this gave systematics a new excitement for him, as well as for others.
Johnson continued to serve on the Council of the Linnean Society of New South Wales, of which he had for two terms been President, and as a member of the Rudi Lemberg Travelling Fellowship Committee of the Australian Academy of Science. He also regularly participated in international congresses, and served on committees of the International Association for Plant Taxonomy and also the International Congress of Systematic and Evolutionary Biology. Whenever possible he took the opportunity of congress excursions to gain wider familiarity with vegetation and floras world-wide. His last, a 'Southern Connections' meeting in January 1997, focusing on Southern Hemisphere biota, was in Valdivia, Chile, with a week-long excursion across the Andes. His final illness was diagnosed a few days after returning from this travel.
To continue effectively in research, Johnson required access to the collections at the Herbarium and contact with colleagues with whom he still had collaborative projects. He became increasingly out of sympathy with trends in management and priorities at the Gardens and made this apparent in often outspoken comments. The Royal Botanic Gardens and Domain Trust expressed its displeasure at these comments but let the arrangement continue since Director Professor Carrick Chambers had championed the value of his continued scientific work. After a change of senior management, however, his status as Honorary Research Associate was terminated in 1996. This 'dishonourable discharge' from the place he had served throughout his career caused him much bitterness although he was encouraged by expressions of support from colleagues within the Royal Botanic Gardens and internationally.
His status as an Associate was restored in early 1997, but by then he had received the diagnosis of brain tumours that had spread from a melanoma removed the previous year. After radiotherapy he made a brief improvement and returned for some weeks to botanical work at the Gardens. He was working on a manuscript on Eucalyptus to within a few hours of his last conscious time. He died at Royal North Shore Hospital, in the Sydney suburb of St Leonards a few days later, on 1August 1997.
It is with the classification of the eucalypts that Johnson is especially associated. Eucalyptus (almost 800 species(4)), together with the allied genera Angophora (15 spp.) and Corymbia (113 spp., segregated from Eucalyptus by Hill and Johnson [118]) form Australia's most prominent tree group. Johnson took an early interest in them because of their importance in characterizing Australian plant communities. Enlisted to help when Bob Anderson was revising his Trees of New South Wales,(5) Johnson soon became the Herbarium's expert on the eucalypts and many other tree groups. Through fieldwork and study of specimens, he realised that there were many species that had not previously been recognized as distinct. These differed from others not only in their morphological features but in distribution and ecology.
Together with Lindsay Pryor from Canberra, Johnson made many field visits, eventually attempting to see as many as possible of the species in their natural habitats. With Pryor's colleague at the Australian National University, Dugal Paton, they visited remote parts of Australia using light aircraft and Dugal's expertise as a pilot with flying experience in the Second World War.
In 1971 Pryor and Johnson commented that 'It is likely that relatively few taxa still await discovery as a result of exploration in botanically little-known areas' [35], but they were proved wrong by their own further studies of existing collections as well as by much further fieldwork. Solely or with colleagues, Johnson was responsible for naming some 180 eucalypt species(4) – mostly after 1971 – and 69 new subspecies, as well as generically reclassifying 107 species; but his conclusions about eucalypts were equally influential above the species level. He sought to characterize the major groups that represent the principal lines of evolution among them. Johnson and Pryor concluded that the eucalypts consisted of some eight major groups which they informally termed subgenera and discussed [35]. This work included a classificatory listing of all the species recognized at the time, and discussions of character states and variation patterns among species. Here they recognised seven informal 'subgenera' and devised a code (comparable to an acronym of four to six letters) to indicate the classification of each species at all ranks. As noted by Brooker, 'This work which divided the genus [Eucalyptus] into seven subgenera, proved to be a milestone in Eucalyptus biology and became the benchmark for all eucalypt scholars.' (6)
The classification in that work was mutually agreed by both authors but depended most heavily on Johnson's work. He did more than anyone else until the late 1990s to distinguish the major evolutionary lineages among the eucalypts [40, 55]. His clarifying these enabled other later researchers, especially those concerned with DNA sequencing, floral development and ecological adaptation, to focus their efforts appropriately. Such subsequent investigations confirmed many relationships deduced from his morphological studies.(7) Pryor took a leading role in pointing out the ecological and distributional differences among the major groups [70], and these have been followed up by further studies by others.(8) In studies of the Myrtaceae and Myrtales, Johnson drew attention to the eucalypt lineage as one of the major ancient evolutionary branches within its large family [80].
Johnson was precise and thorough in nomenclature, orthography and typification at the predominant levels of family, genus and species and served on international committees in these areas. However, he regarded the rigid application of the rule of priority as irrelevant and time-wasting at intermediate levels that derive their significance from circumscription (content) in a particular classification. In this spirit, publishing with Pryor, he set up a formal and precise system at these levels in the classification of the eucalypts, but it was extracodical – not complying with the priority rules of the International Code of Botanical Nomenclature [35]. The Pryor and Johnson extracodical subgenera have been widely used by researchers, but were largely avoided by D.J. and S.M. Carr, with whom Johnson strongly and openly disagreed on several aspects of eucalypt relationships – though Johnson told colleagues that the areas of agreement between them actually far exceeded the areas of dispute. Johnson was disappointed that other botanists clung to those formalities that he regarded as stultifying and that the subgenera were not more widely referred to in publications. Similarly he hoped that the classificatory codes would be widely applied, since he saw them as a simple way to indicate the lineages and affinities. However, he had underestimated the extent to which new species would be recognized in the near future and the classification modified, mostly by himself and colleagues, so that the codes did not always serve as stable summaries of the classification in the way originally intended; he therefore accepted that their introduction had perhaps been somewhat premature. They were, however, very helpful references for use in activities such as herbarium curation.
Disappointed in the extent of use of the subgeneric names, and convinced that the diversity within the eucalypts was greater than within many groups of genera, Johnson for many years considered that the eucalypts should be divided into several separate genera. The prospect of eight genera, or as many as eleven, in place of Eucalyptus alarmed amateur enthusiasts concerned with Australian plants, and popular journals carried articles championing the status quo. Johnson argued to his colleagues and in print [87] that the case for recognizing separate genera, and so changing plant names, should rest on the same types of evidence, whether one was dealing with such a national icon as Eucalyptus or with less charismatic plants such as saltbushes, heaths or rushes, where extensive name-changes created no such outcry.
Eventually Johnson, with his colleague Ken Hill, concluded that the major evolutionary lineages within the eucalypts would be reasonably represented in the classification by the recognition of one new genus, Corymbia, in addition to Eucalyptus and Angophora. In a substantial paper they formally described Corymbia and transferred to it 80 species of bloodwoods and ghost gums, together with 33 newly recognised species [118]. DNA sequencing studies by Udovicic and Ladiges subsequently confirmed that Corymbia is markedly distinct from Eucalyptus and less closely allied to it than to Angophora.(7) Such genetical distinction implies a long separate evolutionary history. Corymbia has been widely adopted by botanists but opinion was not unanimous. Ian Brooker, influential eucalypt specialist at the Australian National Herbarium, CSIRO, Canberra, argued against its adoption(6) but there has been convincing endorsement, based on multiple data sets, of the distinction between Corymbia and Eucalyptus by Ladiges and Udovicic.(9)
In his work on the eucalypts Johnson was greatly assisted by Ken Hill and earlier by Don Blaxell (later Assistant Director Living Collections, Royal Botanic Gardens Sydney, and subsequently Honorary Research Associate, returning to studies on eucalypts with Hill). Expert technical assistance from Leonie Stanberg, and earlier from Lani Retter, was important to the progress of the research. Johnson's strongly held views sometimes contributed to difficult relations with other botanists, especially in the eucalypts, but younger colleagues learnt much from him and he relied on their work, especially when administrative responsibilities greatly limited his research time. As a result of this teamwork, Hill and Blaxell, with assistance from Stanberg, continued after Johnson's death to publish work that had been initiated with him [128, 134, 144].
Studies on eucalypts naturally involved Johnson in questions of how this lineage related to other groups within Myrtaceae. Also, comparisons of inflorescence structures in this and other families made clear to him that some terms, such as panicle, were used to describe a variety of fundamentally different structures. Working with me, and recording observations in simple stylised diagrams, he studied inflorescences throughout a comprehensive range of Myrtaceae. In works by Troll(10) we found a more precise descriptive system than any available in English-language publications. However, our studies on trees and shrubs mostly of warm temperate and tropical regions suggested trends different from those envisaged by Troll. We suggested that undue emphasis on European cool temperate herbs had led Troll and his colleagues to mis-read the direction of some trends.
This work led to a substantial paper on Myrtaceae that included a critique of current inflorescence terminology and introduced new terms to facilitate comparison of inflorescences [61]. We realized that it would be necessary to keep publishing on inflorescences if these new concepts were to be spread more widely. The diversity in Myrtaceae had given an excellent basis for reviewing inflorescence structures, but was not sufficient to promote the general applicability of the new concepts. Some of the main concepts and terminology were taken up and applied, for example by Grimes(11) and Weston, in groups such as Fabaceae and Proteaceae, but pressure of other commitments prevented us from continuing work in interpreting inflorescence structures.
Johnson and I returned to the phylogeny of the Myrtaceae, in a study broadened to cover Myrtales [80], using the laborious procedures of the CLAX programme (see below) in its non-computerized form. This gave some novel results that have since been supported by DNA studies. These include distinguishing a Myrtacean clade including Psiloxylon, Heteropyxis and Myrtaceae within Myrtales and abandoning the subfamilies Myrtoideae and Leptospermoideae. We emphasised the lack of close relationship between the sometimes confused Eugenia and Syzygium, the distinctness and antiquity of the eucalypt lineage, and the polyphyletic nature of the assemblage of genera formerly referred to as the Chamelaucioideae. Not all the relationships postulated have survived in the DNA era, but the hypotheses provided a basis for much subsequent work(12)[125].
As mentioned above, a revision of Casuarina was Johnson's first botanical research project. What was intended as a year's undergraduate study became a lifetime interest, although only intermittently active. As with the eucalypts, his work encompassed the levels of both species and genera. Karen Wilson became the colleague on whom he greatly depended in this work. Alone or with Wilson, he described three new genera and many new species, as well as providing treatments for international compendia, the Flora of Australia and several other Floras [64, 73, 76, 86, 88, 92, 93, 97,135].
Pressures of time when book deadlines conflicted with demanding administrative duties led to the new genera being very briefly described with little stated justification and this slowed their acceptance by other botanists. In this case the common morphological features of the family are so conspicuous that they tend to overshadow the substantial differences between the genera. Johnson's work in clarifying the species taxonomy gave a foundation for studies by Barlow of chromosome numbers and apomixis(13) and work on Casuarinaceae is being continued by Karen Wilson.
Early in his career, Johnson embarked on a revision of Persoonia and realised that relationships at the generic level in Proteaceae were much in need of study. Work on Persoonia proceeded intermittently over a long period, eventually much of it being brought to publication with colleague Peter Weston taking the leading role [101, 117, 122].
Chromosome numbers had been shown by Lancaster and Smith-White(14) to be informative in elucidating evolutionary relationships in the family, so Johnson enlisted my help to determine chromosome numbers of additional genera. This led to a wide-ranging collaboration in addressing the subfamilial, tribal and generic classification and evolutionary relationships. Since Proteaceae is almost confined to the Southern Hemisphere and is represented on almost all southern land masses, clarification of relationships among the genera is highly relevant to the historical biogeography of these lands. At the time of our first publication on this subject [24], in which I took a minor role, geological opinion on plate tectonics was still divided, with a few authoritative voices arguing that continental movement was unproven. Unfortunately, this limited the nature and significance of the biogeographical conclusions that we then presented.
Returning to this subject after a decade [53], the major plate tectonic framework had been established and it was clear that Proteaceae revealed numerous intercontinental links that could be correlated with the ages of separation of the various fragments of Gondwana. Hypotheses were presented of the sequence of diversification of various lineages in relation to opportunities for migration. Since then, the techniques of DNA sequencing have become widely applied to plant phylogeny and findings based on analysis of sequence data for Proteaceae have clarified relationships, superseding some earlier conclusions and revealing additional intercontinental links.(15) Johnson welcomed the new investigations but was gratified that our hypotheses were still serving as a starting point for such new studies decades later.
As in much of Johnson's work at the upper taxonomic levels, these papers on Proteaceae are concise summaries of a great deal of information that was not presented in detail but that was rigorously compiled and critically assessed. To illustrate and describe in detail the floral structures that were dissected or fruits that were sectioned, for example, could have been useful, but it would have vastly increased the work.
In the 1960s Obed Evans, formerly Senior Technician at the Botany Department of the University of Sydney, joined the Herbarium staff to assist with work on the Flora of New South Wales series. Evans worked with Johnson on New South Wales Arecaceae (Palmae), Philydraceae, Lemnaceae, Cyperaceae and Restionaceae [15, 19, 20, 21, 26, 27]. In the very large family Cyperaceae their joint work covered only Eleocharis and some Cyperus species [31, 43], but Johnson continued the study, assisted by Karen Wilson. With increasing expertise, and since Johnson had many projects on hand, Wilson soon took the dominant role and became the Australian authority on much of the Cyperaceae. Meanwhile Johnson had also taken up work on Juncaceae. When he distinguished and described 23 new Australian Juncus species [23, 98, 109], some again questioned whether he was a taxonomic 'splitter', especially since several of the new species frequently occurred together, where disturbance favoured spread of the species and masked ecological distinctions. Others experts on Juncaceae, however, such as Elizabeth Edgar at DSIR Christchurch, New Zealand, were in full agreement with his decisions. Wilson worked with Johnson on Juncaceae also and continues that study.
Johnson and Evans found that stem anatomy in Restionaceae was very informative in distinguishing species and that the species customarily included within Lepyrodia were a disparate group [21]. To gain further information, I joined in with chromosome number determinations. The results were perplexing, so Johnson and I widened the investigation to include the numerous Western Australian species. The study, begun in the early 1960s, was repeatedly set aside in favour of work on Proteaceae and Myrtaceae, but proceeded intermittently and was taken up again in the 1990s. Mostly species distinctions were clear-cut (though many were newly recognised), but settling on an adequate generic classification proved difficult, although it was clear that existing classifications were greatly at variance with relationships among the species. As a result, manuscript names or other informal names were applied and used for up to twenty years(16,) (17) and the patience of other botanists was sorely tested waiting for such names to be formalized. In particular, Cutler, studying the anatomy of Restionaceae at Kew, noted numerous discrepancies between his findings and the existing classification(18) but was largely restrained from making taxonomic changes because the Australian members were under study by Johnson and Briggs. Eventually we finalized a new generic classification of these and some allied groups, under pressure for this to be available for two book projects [132, 136]. By this time botanists at the University of Western Australia and at Kings Park and Botanic Gardens, Perth – John Pate, Kathy Meney and Kingsley Dixon – were studying reproduction, regeneration after fire, and aspects of the physiology of the species(17) and had discovered additional new taxa. Collaboration and fieldwork with these botanists and with Peter Linder from Cape Town, South Africa, proved to be very stimulating and added new dimensions to the study. Especially during field work, Pate and Johnson delighted in matching wits, with puns to the fore. As grass-like plants, lacking conspicuous flowers, and occurring mostly on low-nutrient soils, this had been an extremely neglected group until recent decades. Our studies revealed 54 species that had not been botanically named.
Describing the new genera and necessary changes to the classification of many of the named species followed after Johnson's death [126, 127, 133, 142, 143]. Descriptions of the remaining species that we had jointly distinguished, as well as preparation of a treatment of the family for the Flora of Australia, continue now with further 'Briggs and Johnson' papers expected.
In parallel with the morphological study of Restionaceae, DNA sequencing of representative species was begun, with expert advice from John Thomson and Peter Weston and assistance from Simon Gilmore, and later Adam Marchant and Carolyn Porter. Johnson enthusiastically supported this development, believing that analyses of macromolecular data gave much improved insights into phylogeny. He did not live to see the publication of the main results from this study to date,(19) but was convinced by early findings that the genera Hopkinsia and Lyginia should be excluded from Restionaceae. He therefore shares authorship of the paper establishing the new families Hopkinsiaceae and Lyginiaceae [140], members of the order Poales (as is Restionaceae) and most closely allied to Anarthriaceae; the recognition of these new families has been criticized by Chase et al.(20)
The groups already mentioned were major themes of Johnson's studies but, especially in his early years, he investigated a wide range of plant groups, making extensive and highly informative, but sometimes almost unreadable, annotations on the herbarium folders. In Oleaceae he published a review of the generic classification of world-wide relevance [10]. Indeed a study of the phylogeny and classification of Oleaceae more than four decades later (21) cited Johnson's work as the most recent review of the entire family and supported many of his conclusions when discussing new findings. Some other groups he investigated significantly, but then passed on his findings as notes or in discussion to others who had more prospect of bringing research to fruition. The value of such preliminary work was acknowledged by Bryan Barlow in Loranthaceae and Alma Lee in Xanthorrhoeaceae.
Johnson's accounts of the families of cycads and the Zamiaceae of Australia [12] and of the New South Wales members [13] have been mentioned. Although this was early in his career, he did not hesitate to put forward a new classification on a world-wide basis. He published a review and synopsis of the classification of the families and genera of the Cycadales throughout their range, describing a new family, Stangeriaceae, for a genus of south-east Africa, a new tribe for a genus of Central America and another tribe that included genera in Australia and Africa. Comments were made on the morphological terminology appropriate to this plant group, and the revision of the Australian species of Zamiaceae required the unravelling of complex synonomy. Johnson's work remained of sufficient note that he and colleague Karen Wilson were, decades later, asked to contribute the whole taxonomic treatment of cycads to an international collaborative project [95].
Johnson has been considered by some to be a 'splitter', applying an unduly narrow species concept, but many more Australian Zamiaceae are now recognised than in his treatment.(22) Work on cycads is being continued by his colleague Ken Hill and part, but not all, of the increase in numbers has come with new discoveries resulting from easier access to much of northern Australia.
Johnson's interest in the theory of systematics was demonstrated in his most substantial early publication [12] where he outlined the theoretical foundations that guided his study of cycads world-wide. In subsequent publications he continued to ensure that underlying assumptions were made explicit, although this was not then commonly done by botanical taxonomists.
His main theoretical contribution was in the late 1960s when taxonomy was influenced by new developments that received the name 'numerical taxonomy'. Data bases of the characters of 'operational taxonomic units' were being developed and computer processing gave new options for analysing these data. Numerical taxonomy used similarity measures to build 'trees' of relationships. Johnson entered the scene as a critic of these approaches, with added confidence from studies recently made to gain familiarity with modern mathematical concepts. He concluded that numerical taxonomy was based on false assumptions and published a detailed critique of overall similarity and general-purpose classifications [32]. This was so highly regarded that the prestigious journal Systematic Zoology took the unusual step of reprinting it with a short addendum [34]. Hull, writing more than thirty years later, observed that 'I find his objections to be as well taken now as they were then'.(23) Hull also noted that the pioneers of numerical taxonomy, Sokal and Sneath, 'would have very much liked to persuade Johnson to join with them in their efforts to improve taxonomic principles. They failed. Johnson joined no "school" of taxonomy but tended to his Eucalypts'.
Johnson rejected overall similarity but later saw that there was a better way to determine relationships between organisms, based on the common possession of evolutionary advancements (now known as synapomorphies). He developed this approach independently of the works of Willi Hennig, which he did not read until much later, if at all. He developed an algorithm and procedure for data analysis by grouping first the taxa with the largest number of synapomorphies and progressively adding those with fewer common advanced features. This sequential method, as he claimed, reflected the procedure of systematists building up a structure of relationships from the closely similar to the more dissimilar. This procedure was outlined and used, with laborious manual manipulation, to develop phylogenetic trees for parts of the Proteaceae [53] and the Myrtales and Myrtaceae [80]. His eldest son, Christopher, then a research student in computer science, produced a computerized version of the analysis procedure (termed CLAX), but this was never fully developed. Soon cladistics was becoming widely established, with its theoretical and practical basis developed by others, and Swofford was producing programmes with a range of options that could be selected to match the assumptions on which the analysis was to be based. Johnson's contribution to the demise of numerical phenetics was significant in its time, but CLAX had impact only through the contribution it made to phylogenetic understanding of the groups to which it was applied.
Johnson [90] distinguished between two scientific approaches: 'Some scientists are analysts, strongly influenced by recent philosophies of science and concerned to demonstrate their purity of method, however inadequate the method may be in its coverage of the phenomena of nature. Others are synthesists, less concerned with rigour or the appearance of it, but certainly not less concerned with truth. The latter are interested in forming a picture of what really happens, or happened, in the light of all reasonably reliable evidence that they can bring to bear.' Hull (23) observed that both sorts of scientists are needed if science is to progress and that Johnson clearly saw himself as of the second sort.
Johnson was largely responsible for initiating ecological work in his organisation,(24) with a vegetation mapping programme that provided baseline data about the plant communities of regions surveyed. He also encouraged the Royal Botanic Gardens to submit comments on the major environmental issues of the day, such as the conservation of rainforests, better land management in the Western Division, or recommendations of specific areas to be conserved. Occasionally he had actively to support his staff when they made statements in scientific publications or to the media that displeased senior officers in other government agencies; this arose when scientific conclusions conflicted with short-sighted policies.
Strongly committed to environmental conservation, Johnson was in the vanguard in warning of issues that are only now receiving wide attention. Especially he emphasised the link between weed invasion and altered nutrient status in soils of naturally low fertility, the value of retaining remnants of native vegetation in rural areas, and the importance of safeguarding the detailed regional record of genetic diversity by using local provenances in plantings of native species.
It was characteristic of Johnson, in the Preface to a popular book, Flowers and Plants of New South Wales and Southern Queensland, not to be content with bland remarks but to give a strong message. The opportunity to get that message to a wider audience in a non-technical context was not to be missed:
On the local front, resist by all legal means the unnecessary fouling of gullies by residential or other development at their heads, leading to mineral enrichment and choking by weeds. Resist 'reclaiming' (a profoundly dishonest word) of swamps. Prevent building on headlands and unnecessary artificial revegetation of sand-dunes. Oppose clearing, mowing, planting of roadsides; let the native vegetation or even harmless 'weeds' grow – they will support a rich life of invertebrate animals and some birds and other vertebrates (though certain noxious weeds cannot be tolerated and harbour for rabbits must sometimes be destroyed). Keep even the smallest patches of native or semi-native vegetation – the large reserves alone are not enough. [54]
Another conservation message was primarily addressed to agriculturalists: 'Large areas of the Australian landscape derive much of their character from trees that are survivors from forests or woodlands previously existing in areas now mainly cleared. Many of these trees are already old, and grazing and cultivation are preventing natural establishment of their progeny' [36]. Here Johnson and Briggs argued that natural regeneration also preserves scientifically valuable information about the pre-existing vegetation and emphasised the importance of using locally collected seed in revegetation projects, in order to maintain local genetic provenances.
From the 1950s to the early 1970s, a period when National Parks and Wildlife Service organizations were lacking or embryonic in Australia, there were only a few ecologists with broad knowledge of the flora and vegetation. Systematists had been major champions of nature conservation and served in some roles that would now mostly fall to ecologists. Johnson was influential in this way, especially as an expert member of New South Wales Government committees that reviewed National Parks, State Parks and Reserves in 1967 and subsequently the Committee of Inquiry on Differences and Conflicts between Interests of Parks and Conservation Authorities, Scientific Bodies and Mining Companies (the 'Sim Committee'). His unpublished report on the conservation value of the Kurnell Peninsula was a precursor to the designation of that area as a National Park.(25) He chaired a committee of the New South Wales Government investigating the decline of planted Norfolk Island pines, a striking feature along many coastal beaches. Research commissioned by the committee found that the trees were suffering the effects of pollution by non-biodegradable detergents in wind-borne sea spray. Much later, Johnson provided information on eucalypt diversity to assist colleagues preparing a comprehensive report on the World Heritage values of the Blue Mountains west of Sydney; the area's outstanding diversity of eucalypts was a critical feature emphasised in the successful nomination for its inclusion on the World Heritage List.
Accounts of Johnson's life and achievements by Benson,(24) Hull(23) and Briggs(26) were included in an issue of the journal Telopea dedicated to him in 1996, his 71st year. A short obituary later appeared in Telopea.(27)
He was awarded the Clarke Medal of the Royal Society of New South Wales (1979), and the Mueller Medal of ANZAAS (1984), elected a Fellow of the Australian Academy of Science (1986), and made a Member of the Order of Australia (AM) for services to botanical science (1987), as well as holding honorary or corresponding memberships of the Linnean Society (London), the Botanical Society of America and the American Society of Plant Taxonomists. Two books on very different botanical subjects(17, 28) were dedicated jointly to Johnson and Briggs and, shortly after his death, the preface to papers from a symposium on Proteaceae dedicated the volume 'to all those people who have in one way or another been influenced by Johnson ('Grandfather Proteaceae' as dubbed at the Symposium)'.(29)
Johnson left a legacy in the Royal Botanic Gardens Sydney, which was transformed into a much more vigorous and forward-looking organization during his directorship, with satellite gardens under active development and flourishing education programmes. He was also responsible for the greatly increased breadth and quality of its scientific programmes.
Alone or with colleagues, he distinguished and described four new families of vascular plants, 33 new genera and some 286 species (including posthumous publications), also reclassifying a further 395 others. He has been criticized for being too prompt to alter classifications, thus changing plant names, and not considering sufficiently the confusion resulting for the many non-specialist users of such names. But the insight that prompted the changes, borne of much observation and critical comparison, has mostly led other experts studying the same groups to come to agree with his decisions. Colleagues are continuing to publish work initiated jointly with him; descriptions of thirty new species of Restionaceae, in particular, are yet to be published jointly.
The spectrum of plant families in which Johnson made major clarifications and improvements to classification is remarkably broad, including Casuarinaceae, Myrtaceae, Oleaceae, Proteaceae, Restionaceae and Zamiaceae. In the eucalypts, he and his colleagues delimited evolutionary lineages and systematized available information into a framework that greatly clarified this large and complex group.
In phylogenetics, Johnson's work was done mostly just before a turning point in the subject when powerful cladistic packages and macromolecular data would become widely available. As a result, hypotheses that he developed, solely or jointly, have in some instances been robustly supported but in other cases have been negated by the new data. It is a tribute to the thoroughness of his investigations, the quality of his reasoning and the scope of the questions that he tackled that his phylogenetic hypotheses for the eucalypts, Myrtaceae, Proteaceae, Oleaceae and Myrtales were all still being quoted as starting points for investigations by new approaches many years later.(7, 12,15, 21)
Johnson has been commemorated by other botanists by the naming of the following species: Baumea johnsonii K.L. Wilson (Cyperaceae), Davidsonia johnsonii J.B. Williams & G. Harden (Cunoniaceae), Eucalyptus johnsoniana Brooker & Blaxell (Myrtaceae), Grevillea johnsonii McGill. (Proteaceae), Macrozamia johnsonii D.L. Jones & K.D. Hill (Zamiaceae), Notelaea johnsonii P.S. Green (Oleaceae), Sclerolaena johnsonii (Ising) A.J. Scott (Chenopodiaceae), Typhonium johnsonianum A. Hay & S. Taylor (Araceae) and Xanthorrhoea johnsonii A.T. Lee (Xanthorrhoeaceae).
Johnson never sought research students but worked closely with colleagues in his own institution. Several of those colleagues, especially Karen Wilson, Peter Weston, Ken Hill and I, made careers for some decades there and, as a result, his influence may not have been disseminated as widely around Australia as might have happened if there had been more movement of his colleagues. Nevertheless, his reputation in Australia and internationally was strong and his team in Sydney was productive and notable for the broad synthesis that characterized their studies. In 2001, the Editorial Committee of Australian Systematic Botany determined that the journal would publish a continuing series of invited review articles critically evaluating key areas of systematic botany in Australia. Deciding to name these to commemorate a major contributor to this field, they chose the title, the Johnson Review Series.
Johnson has been seen as unduly vehement, and was certainly demanding of high standards of logical thought, clear expression and wide knowledge. He was often a stern critic but many younger botanists, in particular, referred warmly to kindness and encouragement received from him and his generosity in sharing his knowledge. Especially he supported several botanists whose views were, he believed, being rejected primarily for their unorthodoxy rather than on evidence. A very large number of publications over several decades acknowledged his constructive comment, or discussion with him.
Peter Raven, eminent botanist and Director of the Missouri Botanical Garden, U.S.A., summed up Johnson's international reputation:
Our knowledge of Australian plants has been greatly improved as a result of the industrious, intelligent and forceful career of Lawrie Johnson. He has given us new insight into several of the most important groups of plants in Australia – ones that are leading components in the vegetation, and most interesting biogeographically He was never afraid to take on difficult problems in systematics, and he made important contributions to our understanding of every group that he studied. Few have or could have accomplished so much. (30)
This memoir was originally published in Historical Records of Australian Science, vol.13, no.4, 2001. It was written by Barbara G. Briggs, Honorary Research Associate (formerly Senior Assistant Director Scientific), Royal Botanic Gardens, Sydney.
Numbers in brackets refer to the references, numbers in square brackets refer to the bibliography.
I am grateful for the constructive comments of Merle Johnson, Karen Wilson, Ken Hill, Peter Weston, Peter Wilson and Leonie Stanberg in preparing this account.
(Further joint publications are in preparation by Johnson's former colleagues.)
© 2024 Australian Academy of Science
